What makes a good neighborhood? Interaction of spatial scale and fruit density in the predator satiation dynamics of a masting juniper tree

Spatio-temporal variability in fruit production (masting) has been regarded as a key mechanism to increase plant fitness by reducing seed predation. However, considerably more effort has been devoted into understanding the consequences of temporal rather than spatial variations in fruit crop for plant fitness. In order to simultaneously evaluate both components, we quantify fruit production and pre-dispersal damage by three arthropod species (mites, chalcid wasps and moths) in the Spanish juniper (Juniperus thurifera) during 3 years in a spatially explicit context. Our aims were to assess (1) the interaction between fruit production and pre-dispersal fruit damage by arthropods, (2) the potential interference or competition between arthropods, and (3) the form of the phenotypic selection exerted by arthropods on fruit traits considering the spatial context. Arthropods damaged a substantial fraction of fruits produced by Spanish juniper with levels of damage showing sharp inter-annual variations. Fruit damage by mites was negatively related to yearly fruit crop and positively correlated at individual trees fruiting in consecutive years. Increased interspecific interference was an additional consequence of reduced fruit availability during small crop years. During a masting year, fruit damage by less mobile species such as mites was negatively affected by tree crop size, and no spatial structure was observed for mite damage. The incidence of chalcid wasps was low, so the spatial pattern of seed predation was unclear, and no preferences for fruit or seed traits were detected. Conversely, moths selected larger fruits and their incidence on trees was spatially aggregated up to 20 m, with predation levels being negatively affected by fruit abundance at the patch level, suggesting a positive density-dependent effect of neighbors on fruit output. These results highlight the importance of including the spatial component to understand complex species interactions at local scales.     

Feeding of small rodents on seeds and fruits: a comparative analysis of three species of rodents of the<Emphasis Type="Italic">Araucaria</Emphasis> forest,southern Brazil

We investigated the feeding of three small-sized sigmodontine rodents (Oligoryzomys nigripes Olfers, 1818,Akodon montensis Thomas, 1913, andDelomys dorsalis Hensel, 1872) on fruiting plants in theAraucaria forest of southern Brazil. We offered to wild-captured animals fruits of 22 plant species in 14 families. For each individual, we recorded consumption patterns of pulp and seed. We also analyzed fecal samples to determine the relative abundance of invertebrates, fruit and seed parts, and fungi. The three rodents showed marked potential frugivory, feeding on most species offered. OnlyO. nigripes showed differences in relation to seed-size categories, consuming a high proportion of ‘small’ (diameter ≤2.5 mm) vs ‘large’ seeds. The rodents may be involved in seed dispersion of three plant species, by defecation of viable seeds of twoLeandra (Melastomataceae) species and also by removing the pulp ofMyrceugenia miersiana (Myrtaceae) fruits without swallowing or damaging their relatively large seeds, thus, potentially increasing germination rates ofM. miersiana. Diet analyses indicated no significant difference among proportions of seed/fruit in the fecal samples of the three species of rodents (33 to 41% of plant material). Our results suggest that the three focal species are the main small-mammal seed predators on the ground ofAraucaria forests.     

The impact of consumption of fruit by vertebrate and invertebrate frugivores on the germination success of an Australian rainforest seed

Abstract The consumption of fruits by vertebrates and invertebrates can be both advantageous or detrimental to the survival of the seeds they contain. This study investigated the effect of fruit size and consumption of fruit pulp by rodents and beetles on the germination of the seeds of Acmena graveolens, a tropical rainforest canopy tree found in northern Australia. As fruit size increased, germination success and the amount of pulp remaining on the fruits was greater. When beetles were absent, germination success was highest when most of the pulp was removed by rodents, suggesting that they removed an inhibitor of germination. When beetles were present, germination success did not differ significantly across pulp categories, so beetles apparently enhanced germination in seeds with little pulp initially removed, possibly by further removal of fruit pulp. In this study, both rodents and beetles enhanced germination success of A. graveolens seeds by consumption of fruit pulp. Acting as facilitators of germination is a relatively unusual role for both these frugivores that are generally considered to act as seed predators or (in the case of rodents) dispersers.     

Temporal variation in the effects of habitat fragmentation on reproduction of the Mediterranean shrub <Emphasis Type="Italic">Colutea hispanica</Emphasis>

Habitat fragmentation poses a major threat to the viability of plant populations. However, the intensity of fragmentation effects may vary among years. We studied two possible effects of habitat fragmentation (patch size and isolation) on the reproduction and proportion of damaged fruits in 24 patches of the self-compatible shrub Colutea hispanica for three consecutive years with different climate conditions. We also studied the effect of fragmentation on the incidence of two main pre-dispersal seed predators, the butterflies Iolana iolas and Lampides boeticus. High between-year variability was found in number of viable seeds per fruit, number of fruits per plant, total number of viable seeds per plant and proportion of damaged fruits. In 2003, small, isolated patches had a higher fruit set and number of fruits per plant. The proportion of damaged fruits was significantly lower in isolated populations in 2003, while it was very high in all patches in 2004 and 2005. High between-year variability was also found in the proportion of fruits per plant with I. iolas eggs. In 2003 isolated patches had a lower proportion of fruits with I. iolas eggs, but no significant effect of patch size and isolation was found in 2004 or 2005. The proportion of fruits with L. boeticus eggs was similar in the three years of study, although it was slightly higher in large, non-isolated patches in 2003. Thus, the effects of fragmentation on plant reproduction cannot be generalized from one single-year survey. In contrast to the generally accepted idea that fragmentation reduces plant reproduction, plant fitness may increase in isolated patches in years with high fruit production and low seed predation.     

Allometric allocation in fruit and seed packaging conditions the conflict among selective pressures on seed size

Selective pressures on seed size could vary among the different stages of plant life cycles, so no simple relation could explain a priori its evolution. Here, we determined the relationships between seed size and two fitness components—seed dispersal and survival from predation—in a bird-dispersed tree, Crataegus monogyna. We interpret these relationships in relation to the patterns of mass allocation to fruit and seed components. Selection patterns were assessed at two levels (1) selection pressures on the parent tree; comparing seed dispersal efficiency among individual plants and (2) selection pressures at the individual seed level; comparing seed size variation (i) before and after dispersal, and (ii) before and after postdispersal seed predation. Dispersal efficiency (percentage of seed crop dispersed) was positively correlated with fruit mass and fruit width. Differences in crop size did not offset this effect, and larger seeds were overrepresented in the seed rain relative to the seed pool before dispersal. However, the advantage of larger seeds during the dispersal stage was cancelled later by an opposite selection pressure exerted by seed predators. As a result, smaller seeds had a higher probability of surviving postdispersal seed predation, establishing an evolutionary conflict imposed by the need for dispersal and the danger of being predated. Birds and rodents preferentially selected highly profitable fruits and seeds in terms of the relative proportion of their components. Larger fruits had a higher pulp to seed proportion than smaller ones, and all seeds had the same proportion of coat relative to the embryo-plus-endosperm fraction. Hence, although predator pressures were stronger than disperser ones, larger seeds invested proportionally less in structural defense than in dispersal.     

Modelling oldfield species richness in a mountain area

Seed predation by rodents affects plant population dynamics and it mayrespond to changes in vegetation structure at forest edges. This studyinvestigates the magnitude and direction of a potential edge effect intemperatedeciduous forests, and it seeks possible explanations based on predatorabundance and vegetation structure. The study was conducted at twelve forestedges in northern Switzerland; in six sites all shrubs at the forest edge wereremoved, whereas the other sites maintained a structurally rich shrub layer. Ineach site three transects were laid out from the edge towards the forestinterior. In six plots along these transects (at 1, 3, 6, 12, 25, 50m) we studied seed removal from experimental dishes, rodentabundance in live-traps and four characteristics of the vegetation structure.For both woody seed species (Prunus avium,Viburnum lantana) predation was significantly higher nearthe forest edge in the structurally rich sites; in the other sites no suchgradient was found. Selectively accessible dishes revealed that rodents werethemain predators, whereas predation by insects or molluscs was not observed.Abundance of rodents (Apodemus flavicollis, A.sylvaticus, Clethrionomys glareolus) washighestunder dense shrubs close to the forest edge. In the structurally rich sitesthere was a clear gradient of decreasing shrub cover from the edge towards theforest interior; a weaker gradient was observed in the structurally poor sites.We conclude that high shrub cover near the forest edge is the main determinantfor edge effects in seed predation, and edges without a shrub belt show no sucheffect.     

Fruit removal and postdispersal survivorship in the tropical dry forest shrub Erythroxylum havanense: ecological and evolutionary implications

We studied the relationship between the removal rate and the spatiotemporal availability of ripe fruits of the tropical deciduous shrub Erythroxylum havanense in western Mexico. We also evaluated the effects of dispersal on seed survival during the first stages of establishment. Fast and early dispersal should be favored in E. havanense, since propagules have more time to grow and accumulate resources before the beginning of the severe dry season. In general, high rates of fruit removal imply faster and earlier dispersal. Thus, plants producing large crops should benefit from high removal rates, which will increase the probability of successful establishment by their progeny. To characterize both individual and population fruiting patterns, we made daily counts of fruits on 51 plants arranged in six clumps of different sizes. The daily number of fruits removed per plant was higher for plants with larger initial crop sizes and larger numbers of ripe fruits on a given day, but decreased as clump size increased. Additionally, we monitored postdispersal survival and germination in an experiment manipulating seed density, distance from adult plants, and seed predation. Early establishment was independent of density or distance, and vertebrate seed predation was the main agent of seed mortality. Our results indicate that the critical variable with respect to fruit removal is the number of fruits a plant produces, large plants having higher dispersal rates. Large plants are also more likely to have more seeds escaping postdispersal seed predation.     

Are introduced rats (<Emphasis Type="Italic">Rattus rattus</Emphasis>) both seed predators and dispersers in Hawaii?

Invasive rodents are among the most ubiquitous and problematic species introduced to islands; more than 80% of the world’s island groups have been invaded. Introduced rats (black rat, Rattus rattus; Norway rat, R. norvegicus; Pacific rat, R. exulans) are well known as seed predators but are often overlooked as potential seed dispersers despite their common habit of transporting fruits and seeds prior to consumption. The relative likelihood of seed predation and dispersal by the black rat, which is the most common rat in Hawaiian forest, was tested with field and laboratory experiments. In the field, fruits of eight native and four non-native common woody plant species were arranged individually on the forest floor in four treatments that excluded vertebrates of different sizes. Eleven species had a portion (3–100%) of their fruits removed from vertebrate-accessible treatments, and automated cameras photographed only black rats removing fruit. In the laboratory, black rats were offered fruits of all 12 species to assess consumption and seed fate. Seeds of two species (non-native Clidemia hirta and native Kadua affinis) passed intact through the digestive tracts of rats. Most of the remaining larger-seeded species had their seeds chewed and destroyed, but for several of these, some partly damaged or undamaged seeds survived rat exposure. The combined field and laboratory findings indicate that many interactions between black rats and seeds of native and non-native plants may result in dispersal. Rats are likely to be affecting plant communities through both seed predation and dispersal.     

Weak interactions between avian and insect frugivores: the case of Pistacia terebinthus L. (Anacardiaceae)

Vertebrate frugivores often feed on fruits upon or within which insects also feed, yet little information exists on the potential magnitude of interactions between these consumers. The Mediterranean shrub Pistacia terebinthus, the birds that consume its fruits, and the wasps that feed upon its seeds are examined in this study. P. terebinthus produces a highly variable fraction of final-sized red fruits that never become mature (green-colored). Red fruits can be immature, parthenocarpic, aborted, or attacked by wasps, and their pulp is much less nutritious than that of mature fruits. A total of 20 bird species consumed the fruits in the study area. Legitimate dispersers accounted for 39% of the total fruit removal, while pulp eaters and seed predators accounted for the remainder. Birds strongly preferred the mature fruits (only 4% of the fruits consumed were red). The incidence of wasps in the seeds ranged from 0 to 42% of the crop in 1989 and from 0 to 24% in 1990. The influence of avian and insect frugivore guilds on each other appears to be quite low because of the narrow overlap in resource utilization by birds and wasps, and an overall low intensity of wasp seed predation. From an evolutionary perspective, the possible ability of wasps to preclude fruit maturation appears not to be attributable to the present interaction with avian frugivores.     

Taste and Physiological Responses to Glucosinolates: Seed Predator versus Seed Disperser

In contrast to most other plant tissues, fleshy fruits are meant to be eaten in order to facilitate seed dispersal. Although fleshy fruits attract consumers, they may also contain toxic secondary metabolites. However, studies that link the effect of fruit toxins with seed dispersal and predation are scarce. Glucosinolates (GLSs) are a family of bitter-tasting compounds. The fleshy fruit pulp of Ochradenus baccatus was previously found to harbor high concentrations of GLSs, whereas the myrosinase enzyme, which breaks down GLSs to produce foul tasting chemicals, was found only in the seeds. Here we show the differential behavioral and physiological responses of three rodent species to high dose (80%) Ochradenus’ fruits diets. Acomys russatus, a predator of Ochradenus’ seeds, was the least sensitive to the taste of the fruit and the only rodent to exhibit taste-related physiological adaptations to deal with the fruits’ toxins. In contrast, Acomys cahirinus, an Ochradenus seed disperser, was more sensitive to a diet containing the hydrolyzed products of the GLSs. A third rodent (Mus musculus) was deterred from Ochradenus fruits consumption by the GLSs and their hydrolyzed products. We were able to alter M. musculus avoidance of whole fruit consumption by soaking Ochradenus fruits in a water solution containing 1% adenosine monophosphate, which blocks the bitter taste receptor in mice. The observed differential responses of these three rodent species may be due to evolutionary pressures that have enhanced or reduced their sensitivity to the taste of GLSs.     

Friend or foe? Disparate plant–animal interactions of two congeneric rodents

Food and water resources are limiting factors for animals in desert ecosystems. Fleshy fruits are a rare water source in deserts and when available they tend to attract a wide variety of organisms. Here we show that two congeneric rodent species, Acomys cahirinus and A. russatus, employ different fruit eating strategies that result in either dispersal or predation of the small seeds of the desert plant Ochradenus baccatus. The nocturnal A. cahirinus leaves intact seeds when consuming O. baccatus fruits and thus, acts mainly as a seed disperser; whereas the diurnal A. russatus consumes the whole fruit and digests the seeds and thus, acts mainly as a seed predator. Acomys russatus is subjected to the toxic products of the glucosinolates-myrosinase system found in O. baccatus fruits. Acomys cahirinus avoids the toxic compounds by consuming the pulp only, which contains glucosinolates but not the seeds that contain the enzyme that activates them. We suggest that the behavioral responses exhibited by A. russatus are the result of physiological adaptations to whole fruit consumption that are absent in A. cahirinus. Our results shed new light on the ecological divergence of the two congeneric species.     

Effect of mango weevil (Coleoptera: Curculionidae) damage on mango seed viability in Hawaii

The mango weevil, Cryptorhynchus (= Sternochetus) mangiferae (F.), is a federally quarantined pest that prevents shipment of mangos from Hawaii into the continental United States. Although this monophagous weevil allegedly causes reduced seed germination, damage to the fruit pulp, and premature fruit drop in mangos, there are few studies examining these potential sources of crop loss. We conducted studies to assess the effect of mango weevil infestation on seed viability while making observations on the frequency of pulp feeding. Naturally infested seeds from mature fruit were planted in pots and scored for successful germination. Germination rates for infested seeds were equal to those of uninfested control seeds in a polyembryonic cultivar ('Common'), whereas germination was significantly reduced for infested seeds of a monoembryonic cultivar ('Haden') compared with uninfested control seeds but germination of infested seeds was still > 70%. To assess seed tolerance of damage, seeds were artificially damaged by cutting away 25, 50, or 75% of the cotyledon before planting and scored for germination. None of the damage treatments was significantly different from the undamaged controls, indicating that mango seeds can withstand substantial damage and still germinate successfully. Over the 2-yr period we conducted experiments, only four of 3,602 mango fruits (0.11%) showed evidence of direct feeding damage to the pulp. Results suggest that C. mangiferae is a less serious pest of mangos than previously thought.     

Temporal Variation in Seed Predation by Insects in a Population of Syagrus romanzoffiana (Arecaceae) in Santa Catarina Island, SC, Brazil

Insect seed predation may vary depending on seed production. The present study considers the hypothesis that the rates of seed predation tend to be smaller in years of higher fruit production. Thus, we monitored the production of fruits and predation of seeds of the palm Syagrus romanzoffiana over 2?years in the Atlantic Forest (Parque Municipal da Lagoa do Peri, Florianópolis, SC, Brazil), between July 2006 and June 2008. Plots of 0.25?m² were fitted under 20 mother plants and fruits were monthly collected for assessment of abundance and seed predation. There was variation in fruit production between the 2?years and among reproductive plants. Predation rates were high and occurred in the predispersal phase by the Curculionidae Revena rubiginosa Boheman, Anchylorhynchus aegrotus Fahraeus, and Anchylorhynchus variabilis Gyllenhal. Seed predation by these species of Anchylorhynchus is first registered in the present study. In average, about 60% of the seeds monthly produced in the population tend to escape insect predation in year of high or low production, becoming available for recruitment. The predation rate was not related to the amount of fruits produced per reproductive plant. Also, different than expected, there was a positive relation between the rates of seed predation and the total of fruits produced monthly on the plots. Thus, no evidence for the satiation of insect seed predators was found in this study with S. romanzoffiana.     

Bottom-up and top-down effects in a pre-dispersal seed predation system: are non-predated seeds damaged?

Seed predators that severely affect seed germination rates are well known for many plant species. Here, we hypothesised that due to differences in resource allocation within fruits, seed predation can negatively affect non-predated seeds in infested fruits when predation occurs during fruit maturation (a ‘top-down’ effect). We addressed this question using a system of bruchid beetles on Mimosa trees and we also investigated whether seed quality (nitrogen concentration) affects beetle body mass, which would have implications for adult fitness (‘bottom-up’ effect). To assess spatial variation, bottom-up and top-down effects were investigated in two plant populations. Nitrogen concentration was significantly higher in seeds from non-infested fruits than from infested fruits. This supports the hypothesis that resource allocation may differ between seeds from infested and non-infested fruits. Germination experiments showed that seeds from non-infested fruits germinated better than non-predated seeds from infested fruits. It was also confirmed that seed quality affected bruchid body mass. There was also evidence that more resources were taken from well-developed seeds. These results showed that seed predation can damage non-predated seeds.     

Fruit removal efficiency and success: influence of crop size in a neotropical treelet

The reproductive success of animal-dispersed plants is closely linked to the number of seeds that they are able to disperse. The fruit crop size hypothesis states that a plant with large fruit crop size will attract more dispersers than a plant with a smaller fruit crop, which may result in more seeds being dispersed from the foremost. In this study, we experimentally examined the effect of crop size and other factors on primary seed dispersal in a neotropical shrub/tree, Casearia corymbosa (Flacourtiaceae). We used two predictive variables of reproductive success, which produce an accurate picture of seed dispersal ratio: fruit removal efficiency (proportion of a fruit crop removed by frugivores) and fruit removal success (relative contribution of each individual tree to the number of fruits removed in the population). We established two levels of fruit crop size at the C. corymbosa individuals, using plants with large (150 fruits) and small crops (50 fruits). We found that individual plants with larger crops had significantly higher values of fruit removal efficiency (92.6%) and success (5%) than plants with smaller crops (69.3% and 1.3%, respectively). Fruit removal efficiency was related to vegetation type, plant height and fruit width, but the variance explained by these variables was low ( < 8%). Fruit removal success was significantly related to crop size ( > 90% of the variance explained). These results suggest that fruit removal efficiency and success are strongly related to fruit crop size of C. corymbosa plants.     

Do frugivorous birds concentrate their foraging activities on those alien plants with the most abundant and nutritious fruits in the South African Mediterranean-climate region?

Alien plants with abundant and nutritious fruits may compete more effectively for avian dispersal services than native shrubs. This premise was examined by comparing daily foraging activity (visitation frequency, foraging bird number and foraging period) by four frugivorous bird species of different size on fruits of two native and two alien shrub species co-occurring at four different sites. Also, the quantities of seeds consumed daily by each of the four birds species from fruits of the four shrub species were measured and compared with the numbers of fruits and seeds, and the mass of fruits present in the shrub canopies as well as with their fruit monosaccharide concentrations. The quantities of seed consumed daily by four different size bird species (Columba arquatrix, Colius striatus, Pycnonotus capensis, Zosterops pallidus subsp capensis) were positively correlated with the numbers of seeds per m² of canopy area and with fruit mass and fruit monosaccharide content per m² of canopy area, as well as with the monosaccharide concentration of individual fruits, except in the C. arquatrix (African olive pigeon). All four bird species displayed the highest daily visitation frequencies on fruits of the alien Solanum mauritianum which were more abundant and nutritious than fruits of the other alien Lantana camara and fruits the natives Olea europaea subsp africana and Chrysanthemoides monilifera. They also all consumed greater quantities of seed daily from fruits of the alien S. mauritianum than from fruits of the other shrub species. These results corroborate proposals that frugivorous birds concentrate their foraging activities on those alien plants with the most abundant and nutritious fruits.     

Large-scale rodent control reduces pre- and post-dispersal seed predation of the endangered Hawaiian lobeliad, Cyanea superba subsp. superba (Campanulaceae)

Large-scale rodent control can help to manage endangered species that are vulnerable to invasive rodent consumption. A 26?ha rodent snap-trap grid was installed in montane forest on Oahu Island, Hawaii, in order to protect endangered snails and plants. To assess the effectiveness of this trapping operation in reducing fruit consumption and seed predation of the endangered Hawaiian lobeliad, Cyanea superba subsp. superba, pre- and post-dispersal C. superba fruit consumption were monitored for 36 plants at the site with rodent control (Kahanahaiki) and 42 plants at an adjacent site without rodent control (Pahole). Over 47?% of all monitored fruit were eaten on the plants at Pahole compared to 4?% at Kahanahaiki. Images captured using motion-sensing cameras suggest that black rats (Rattus rattus) were the only pre-dispersal fruit consumers. To quantify post-dispersal fruit consumption, and to identify the culprit frugivore(s), mature fruit were placed in tracking tunnels positioned on the forest floor and checked daily. At Pahole, all of the fruit were consumed by rats compared to 29?% at Kahanahaiki. Lastly, to determine if rodents from the sites were predators or dispersers of C. superba seed, fruit were fed to captive black rats and house mice (Mus musculus). Black rats consumed entire fruit, killing all the seed, while mice did little damage to the fruit and seed. Therefore, large-scale rat trapping can directly benefit the reproduction of C. superba subsp. superba. Controlling black rats at restoration sites appears integral to the successful restoration of this endangered plant species.     

Cloning capacity helps seeds of Garcinia xanthochymus counter animal predation

Seed predators have the potential to act as agents of natural selection that influence seed traits and seed fates, which in turn affect the whole plant population dynamic. Accordingly, plants deploy a variety of mechanisms (e.g., resistance and tolerance strategies) to lessen the impact of predation on seed crop or on an individual seed. In this study, we described a novel mechanism, seed cloning strategy, in a tropical plant species in countering animal predation. By conducting field‐ and laboratory‐based germination experiments, we found that both rodent damaged and artificially damaged seed fragments of a large‐seeded tree Garcinia xanthochymus (Clusiaceae) could successfully germinate and establish as seedlings. Tissue culture experiments revealed that G. xanthochymus has no endosperm in seeds, and its seed fragments own strong capacity of differentiation and cloning. Seed damage negatively affected seedling growth and germination, but the seed germination rate was remarkably high. Our study suggests that, seed cloning capacity, adopted by the large‐seeded tree G. xanthochymus may act as a novel strategy counteract for seed predation and would play a significant role in stabilizing the mutualism between plant and animals.     

Assessing ant seed predation in threatened plants: a case study

Erodium paularense is a threatened plant species that is subject to seed predation by the granivorous ant Messor capitatus. In this paper we assessed the intensity and pattern of ant seed predation and looked for possible adaptive strategies at the seed and plant levels to cope with this predation. Seed predation was estimated in 1997 and 1998 at the population level by comparing total seed production and ant consumption, assessed by counting seed hulls in refuse piles. According to this method, ant seed predation ranged between 18% and 28%. A more detailed and direct assessment conducted in 1997 raised this estimate to 43%. In this assessment spatial and temporal patterns of seed predation by ants were studied by mapping all nest entrances in the studied area and marking the mature fruits of 109 reproductive plants with a specific colour code throughout the seed dispersal period. Intact fruit coats were later recovered from the refuse piles, and their mother plants and time of dispersal were identified. Seeds dispersed at the end of the dispersal period had a greater probability of escaping from ant seed predation. Similarly, in plants with late dispersal a greater percentage of seeds escaped from ant predation. Optimum dispersal time coincided with the maximum activity of granivorous ants because, at this time, ants focused their harvest on other plant species of the community. It was also observed that within-individual seed dispersal asynchrony minimised seed predation. From a conservation perspective, results show that the granivorous ant–plant interaction cannot be assessed in isolation and that the intensity of its effects basically depends on the seed dispersal pattern of the other members of the plant community. Furthermore, this threat must be assessed by considering the overall situation of the target population. Thus, in E. paularense, the strong limitation of safe-sites for seedling establishment reduces the importance of seed predation.     

REPRODUCTIVE ATTRITION IN THE RARE CHAPARRAL SHRUB FREMONTODENDRON DECUMBENS LLOYD (STERCULIACEAE)

We examined reproductive attrition in Fremontodendron decumbens to characterize sexual reproduction in this rare California shrub. Reproductive individuals produced an average of 2,900 flower buds in a season, with no significant difference in bud production between two seasons. Because of intense insect predation, <;2% of initiated flower buds became mature fruits. A threefold decrease in predation of flower buds between seasons resulted in an increase in seed output the second season, indicating that seed production was partially predator-limited. Most seeds (97.8%) were dormant due to an impermeable seed coat. Breaking of the coat, mechanically or by heat, allowed high levels of germination. Chamise charate and ash added to the potting medium resulted in the highest level of germination and emergence. Rodents were more important than birds as seed predators, destroying 90% of seeds under parent shrub canopies within 8–10 months. Seeds already integrated into the seed bank were comparatively safe from predation, relative to newly added seeds. If predation was prevented, seeds were long-lived under field conditions (>;80% survived after 5.75 years). Most seedlings produced in unburned chaparral by planting heat-treated seeds in openings between shrubs were destroyed by predators (rodents and insects). All seedlings that escaped predation died during the summer drought. We concluded that sexual reproduction was limited by (in order of importance): 1) lack of fire, 2) predehiscence predation by insects, and 3) postdehiscence predation by rodents. Size distributions from two populations revealed that, despite the apparent absence of sexual reproduction in unbumed chaparral, two unbumed sites contained a large proportion of individuals in small size classes. Excavation of several small individuals demonstrated they were sprouts from the roots of nearby larger shrubs. Because asexual reproduction by rootsprouting circumvents the high attrition of sexual reproductive effort on unbumed sites, rootsprouting may be a significant reproductive strategy of some ‘sprouter’ species in chaparral.